Sensitivity to Interaural Correlation of Single Neurons in the Inferior Colliculusof Guinea Pigs

Sensitivity to changes in the interaural correlation of 50-ms bursts of narrowband or broadband noise was measured in single neurons in the inferior colliculus of urethane-anaesthetized guinea pigs. Rate vs. interaural correlation functions (rICFs) were measured using two methods. These methods compensated in different ways for the inherent variance in interaural correlation between tokens with the same expected correlation. The shape of all rICFs could be best described by power functions allowing them to be summarized by two parameters. Most rICFs were best fit by a power below 2, indicating that they were only slightly nonlinear. However, there were a few fitted functions that had a power of 3–6, indicating marked curvature. Modeling results indicate that the nonlinearity of the majority of rICFs was explicable in terms of the monaural transduction stages; however, some of the rICFs with power greater than 2 require either multiple inputs to the coincidence detector or additional nonlinearities to be included in the model. Discrimination thresholds were estimated at reference correlations of −1, 0, and +1 using receiver operating characteristic (ROC) analysis of the spike-count distribution at each correlation. Thresholds spanned the full possible range, from a minimum of 0.1 to the maximum possible of 2. Thresholds were generally highest with a reference correlation of −1, intermediate with a reference of 0, and lowest with a reference correlation of +1. Thresholds were lowest for the most steeply sloped rICFs, but thresholds were not strongly correlated to the spike rate variance. The lowest thresholds occurred using narrowband noise that was compensated for internal delays, but they were still about three times larger than human psychophysical thresholds measured using similar stimuli. The data suggest that, unlike pure tone interaural time difference, discrimination of a population measure is required to account for behavioral interaural correlation discrimination performance.     

Behavioral Sensitivity to Broadband Binaural Localization Cues in the Ferret

Although the ferret has become an important model species for studying both fundamental and clinical aspects of spatial hearing, previous behavioral work has focused on studies of sound localization and spatial release from masking in the free field. This makes it difficult to tease apart the role played by different spatial cues. In humans and other species, interaural time differences (ITDs) and interaural level differences (ILDs) play a critical role in sound localization in the azimuthal plane and also facilitate sound source separation in noisy environments. In this study, we used a range of broadband noise stimuli presented via customized earphones to measure ITD and ILD sensitivity in the ferret. Our behavioral data show that ferrets are extremely sensitive to changes in either binaural cue, with levels of performance approximating that found in humans. The measured thresholds were relatively stable despite extensive and prolonged (>16 weeks) testing on ITD and ILD tasks with broadband stimuli. For both cues, sensitivity was reduced at shorter durations. In addition, subtle effects of changing the stimulus envelope were observed on ITD, but not ILD, thresholds. Sensitivity to these cues also differed in other ways. Whereas ILD sensitivity was unaffected by changes in average binaural level or interaural correlation, the same manipulations produced much larger effects on ITD sensitivity, with thresholds declining when either of these parameters was reduced. The binaural sensitivity measured in this study can largely account for the ability of ferrets to localize broadband stimuli in the azimuthal plane. Our results are also broadly consistent with data from humans and confirm the ferret as an excellent experimental model for studying spatial hearing.     

Limitations on Monaural and Binaural Temporal Processing in Bilateral Cochlear Implant Listeners

Monaural rate discrimination and binaural interaural time difference (ITD) discrimination were studied as functions of pulse rate in a group of bilaterally implanted cochlear implant users. Stimuli for the rate discrimination task were pulse trains presented to one electrode, which could be in the apical, middle, or basal part of the array, and in either the left or the right ear. In each two-interval trial, the standard stimulus had a rate of 100, 200, 300, or 500 pulses per second and the signal stimulus had a rate 35 % higher. ITD discrimination between pitch-matched electrode pairs was measured for the same standard rates as in the rate discrimination task and with an ITD of +/− 500 μs. Sensitivity (d′) on both tasks decreased with increasing rate, as has been reported previously. This study tested the hypothesis that deterioration in performance at high rates occurs for the two tasks due to a common neural basis, specific to the stimulation of each electrode. Results show that ITD scores for different pairs of electrodes correlated with the lower rate discrimination scores for those two electrodes. Statistical analysis, which partialed out overall differences between listeners, electrodes, and rates, supports the hypothesis that monaural and binaural temporal processing limitations are at least partly due to a common mechanism.     

Behavioral sensitivity to interaural time differences in the rabbit

An important cue for sound localization and separation of signals from noise is the interaural time difference (ITD). Humans are able to localize sounds within 1-2 degrees and can detect very small changes in the ITD (10-20micros). In contrast, many animals localize sounds with less precision than humans. Rabbits, for example, have sound localization thresholds of approximately 22 degrees . There is only limited information about behavioral ITD discrimination in animals with poor sound localization acuity that are typically used for the neural recordings. For this study, we measured behavioral discrimination of ITDs in the rabbit for a range of reference ITDs from 0 to +/-300micros. The behavioral task was conditioned avoidance and the stimulus was band-limited noise (500-1500Hz). Across animals, the average discrimination threshold was 50-60micros for reference ITDs of 0 to +/-200micros. There was no trend in the thresholds across this range of reference ITDs. For a reference ITD of +/-300micros, which is near the limit of the physiological window defined by the head width in this species, the discrimination threshold increased to approximately 100micros. The ITD discrimination in rabbits less acute than in cats, which have a similar head size. This result supports the suggestion that ITD discrimination, like sound localization [see Heffner, 1997. Acta Otolaryngol. 532 (Suppl.), 46-53] is determined by factors other than head size.     

Detection of interaural correlation by neurons in the superior olivary complex, inferior colliculus and auditory cortex of the unanesthetized rabbit

A critical binaural cue important for sound localization and detection of signals in noise is the interaural time difference (ITD), or difference in the time of arrival of sounds at each ear. The ITD can be determined by cross-correlating the sounds at the two ears and finding the ITD where the correlation is maximal. The amount of interaural correlation is affected by properties of spaces and can therefore be used to assess spatial attributes. To examine the neural basis for sensitivity to the overall level of the interaural correlation, we identified subcollicular neurons and neurons in the inferior colliculus (IC) and auditory cortex of unanesthetized rabbits that were sensitive to ITDs and examined their responses as the interaural correlation was varied. Neurons at each brain level could show linear or non-linear responses to changes in interaural correlation. The direction of the non-linearities in most neurons was to increase the slope of the response change for correlations near 1.0. The proportion of neurons with non-linear responses was similar in subcollicular and IC neurons but increased in the auditory cortex. Non-linear response functions to interaural correlation were not related to the type of response as determined by the tuning to ITDs across frequencies. The responses to interaural correlation were also not related to the frequency tuning of the neuron, unlike the responses to ITD, which broadens for neurons tuned to lower frequencies. The neural discriminibility of the ITD using frozen noise in the best neurons was similar to the behavioral acuity in humans at a reference correlation of 1.0. However, for other reference ITDs the neural discriminibility was more linear and generally better than the human discriminibility of the interaural correlation, suggesting that stimulus rather than neural variability is the basis for the decline in human performance at lower levels of interaural correlation.     

Interaural time and level difference thresholds for acoustically presented signals in post-lingually deafened adults fitted with bilateral cochlear implants using CIS+ processing

OBJECTIVES: The main purpose of the study was to measure thresholds for interaural time differences (ITDs) and interaural level differences (ILDs) for acoustically presented noise signals in adults with bilateral cochlear implants (CIs). A secondary purpose was to assess the correlation between the ILD and ITD thresholds and error scores in a horizontal-plane localization task, to test the hypothesis that localization by individuals with bilateral implants is mediated by the processing of ILD cues. DESIGN: Eleven adults, all postlingually deafened and all bilaterally fitted with MED-EL COMBI 40+ CIs, were tested in ITD and ILD discrimination tasks in which signals were presented acoustically through headphones that fit over their two devices. The stimulus was a 200-msec burst of Gaussian noise bandpass filtered from 100 to 4000 Hz. A two-interval forced-choice adaptive procedure was used in which the subject had to respond on each trial whether the lateral positions of the two sound images (with the interaural difference favoring the left and right sides in the two intervals) moved from left-to-right or right-to-left. RESULTS: In agreement with previously reported data, ITD thresholds for the subjects with bilateral implants were poor. The best threshold was approximately 400 microsec, and only five of 11 subjects tested achieved thresholds <1000 microsec. In contrast, ILD thresholds were relatively good; mean threshold was 3.8 dB with the initial compression circuit on the implant devices activated and 1.9 dB with the compression deactivated. The ILD and ITD thresholds were higher than previously reported thresholds obtained with direct electrical stimulation (generally, <1.0 dB and 100 to 200 microsec, respectively). When the data from two outlying subjects were omitted, ILD thresholds were highly correlated with total error score in a horizontal-plane localization task, computed for sources near midline (r = 0.87, p < 0.01). CONCLUSIONS: The higher ILD and ITD thresholds obtained in this study with acoustically presented signals (when compared with prior data with direct electrical stimulation) can be attributed-at least partially-to the signal processing carried out by the CI in the former case. The processing strategy effectively leaves only envelope information as a basis for ITD discrimination, which, for the acoustically presented noise stimuli, is mainly coded in the onset information. The operation of the compression circuit reduces the ILDs in the signal, leading to elevated ILD thresholds for the acoustically presented signals in this condition. The large magnitude of the ITD thresholds indicates that ITDs could not have contributed to the performance in the horizontal-plane localization task. Overall, the results suggest that for subjects using bilateral implants, localization of noise signals is mediated entirely by ILD cues, with little or no contribution from ITD information.     

Bone-conducted sound lateralization of interaural time difference and interaural intensity difference in children and a young adult with bilateral microtia and atresia of the ears

Bone-conducted sound lateralization tests to determine interaural time difference (ITD) and interaural intensity difference (IID) were conducted by means of a self-recording apparatus in 20 children and a young adult with bilateral microtia and atresia of the ear. This apparatus changes ITD automatically from 0 to 2,000 micros at 50 micros/s and IID from 1 to 40 dB at 1 dB's. When ITD exceeds approximately 200 micros/s and IID exceeds 5 dB in normal subjects the sounds are recognized separately. The test stimulus was a continuous narrow-band noise at 500 Hz and 30 dB SL applied to the right and left mastoids through bone vibrators. In the patients with bilateral atresia of the ears, ITD results revealed approximately normal thresholds of discrimination in half the patients and IID results revealed threshold elevation in only 10%. It is noted that bone-conducted sound lateralization abilities of ITD or IID are maintained in many of these patients.     

Detection of Large Interaural Delays and Its Implication for Models of Binaural Interaction

The interaural time difference (ITD) is a major cue to sound localization along the horizontal plane. The maximum natural ITD occurs when a sound source is positioned opposite to one ear. We examined the ability of owls and humans to detect large ITDs in sounds presented through headphones. Stimuli consisted of either broad or narrow bands of Gaussian noise, 100 ms in duration. Using headphones allowed presentation of ITDs that are greater than the maximum natural ITD. Owls were able to discriminate a sound leading to the left ear from one leading to the right ear, for ITDs that are 5 times the maximum natural delay. Neural recordings from optic-tectum neurons, however, show that best ITDs are usually well within the natural range and are never as large as ITDs that are behaviorally discriminable. A model of binaural cross-correlation with short delay lines is shown to explain behavioral detection of large ITDs. The model uses curved trajectories of a cross-correlation pattern as the basis for detection. These trajectories represent side peaks of neural ITD-tuning curves and successfully predict localization reversals by both owls and human subjects.     

Auditory evoked magnetic fields in relation to interaural time delay and interaural correlation

The detection of interaural time differences (ITD) for sound localization depends on the similarity between the left and right ear signals, namely interaural correlation (IAC). Human localization performance deteriorates with decreasing IACs. In order to examine activity related to localization performance in the human cortex, auditory evoked magnetic fields to the ITD of bandpass noises with different IACs were analyzed. When the IAC was 0.95, the N1m amplitudes, i.e., the estimated equivalent current dipole moments, increased with increasing ITD. However the effect of ITD on the N1m amplitudes was not significant when the IAC was 0.5. When the ITD was 0.7 ms, the N1m amplitudes decreased with decreasing IACs. There were no systematic changes in the source location of N1m in the auditory cortex related to changes in ITD or IAC. The results suggest that localization performance is reflected in N1m amplitudes.     

Sensitivity to Interaural Time Differences with Combined Cochlear Implant and Acoustic Stimulation

The interaural time difference (ITD) is an important cue to localize sound sources. Sensitivity to ITD was measured in eight users of a cochlear implant (CI) in the one ear and a hearing aid (HA) in the other severely impaired ear. The stimulus consisted of an electric pulse train of 100 pps and an acoustic filtered click train. Just-noticeable differences (JNDs) in ITD were measured using a lateralization paradigm. Four subjects exhibited median JNDs in ITD of 156, 341, 254, and 91 μs; the other subjects could not lateralize the stimuli consistently. Only the subjects who could lateralize had average acoustic hearing thresholds at 1,000 and 2,000 Hz better than 100-dB SPL. The electric signal had to be delayed by 1.5 ms to achieve synchronous stimulation at the auditory nerves.

Responses of Auditory Nerve and Anteroventral Cochlear Nucleus Fibers to Broadband and Narrowband Noise: Implications for the Sensitivity to Interaural Delays

The quality of temporal coding of sound waveforms in the monaural afferents that converge on binaural neurons in the brainstem limits the sensitivity to temporal differences at the two ears. The anteroventral cochlear nucleus (AVCN) houses the cells that project to the binaural nuclei, which are known to have enhanced temporal coding of low-frequency sounds relative to auditory nerve (AN) fibers. We applied a coincidence analysis within the framework of detection theory to investigate the extent to which AVCN processing affects interaural time delay (ITD) sensitivity. Using monaural spike trains to a 1-s broadband or narrowband noise token, we emulated the binaural task of ITD discrimination and calculated just noticeable differences (jnds). The ITD jnds derived from AVCN neurons were lower than those derived from AN fibers, showing that the enhanced temporal coding in the AVCN improves binaural sensitivity to ITDs. AVCN processing also increased the dynamic range of ITD sensitivity and changed the shape of the frequency dependence of ITD sensitivity. Bandwidth dependence of ITD jnds from AN as well as AVCN fibers agreed with psychophysical data. These findings demonstrate that monaural preprocessing in the AVCN improves the temporal code in a way that is beneficial for binaural processing and may be crucial in achieving the exquisite sensitivity to ITDs observed in binaural pathways.     

A population study of the precedence effect

Data are reported from a population of untrained individuals under lag- and single-click conditions in a discrimination suppression precedence-effect task. The cue to be discriminated was an interaural level-difference (ILD). Each of 91 observers completed 10 runs in a two-interval forced-choice design under a lag-click condition and three runs under a single-click condition. Stimuli were 125-micros rectangular pulses and the interclick interval was 2 ms. Observers were randomly assigned to three groups of approximately 30. Each group was then tested at one stimulus intensity (43, 58, or 73 dB). Mean threshold within each group was greater than 15 dB for the lag-click condition and 6 dB for the single-click condition, although there was substantial interobserver variability. In contrast to [J. Acoust. Soc. Am. 114 (2003) 420] who reported a strong effect of intensity on lag-click ITD discrimination, no effect of intensity was observed on lag-click ILD thresholds. Analysis of over 50,000 near-threshold trials from 302 observers pooled across studies showed a spatial asymmetry in response patterns and a small, but statistically significant effect of gender. A model is proposed which shows that decay of sensory memory and increases in auditory filter bandwidths with intensity may predict the different findings for ILD versus ITD lag-click thresholds.     

The Interaural Time Difference Pathway: a Comparison of Spectral Bandwidth and Correlation Sensitivity at Three Anatomical Levels

Temporal differences between the two ears are critical for spatial hearing. They can be described along axes of interaural time difference (ITD) and interaural correlation, and their processing starts in the brainstem with the convergence of monaural pathways which are tuned in frequency and which carry temporal information. In previous studies, we examined the bandwidth (BW) of frequency tuning at two stages: the auditory nerve (AN) and inferior colliculus (IC), and showed that BW depends on characteristic frequency (CF) but that there is no difference in the mean BW of these two structures when measured in a binaural, temporal framework. This suggested that there is little frequency convergence in the ITD pathway between AN and IC and that frequency selectivity determined by the cochlear filter is preserved up to the IC. Unexpectedly, we found that AN and IC neurons can be similar in CF and BW, yet responses to changes in interaural correlation in the IC were different than expected from coincidence patterns (“pseudo-binaural” responses) in the AN. To better understand this, we here examine the responses of bushy cells, which provide monaural inputs to binaural neurons. Using broadband noise, we measured BW and correlation sensitivity in the cat trapezoid body (TB), which contains the axons of bushy cells. This allowed us to compare these two metrics at three stages in the ITD pathway. We found that BWs in the TB are similar to those in the AN and IC. However, TB neurons were found to be more sensitive to changes in stimulus correlation than AN or IC neurons. This is consistent with findings that show that TB fibers are more temporally precise than AN fibers, but is surprising because it suggests that the temporal information available monaurally is not fully exploited binaurally.     

The choice of stimulation strategy affects the ability to detect pure tone inter-aural time differences in children with early bilateral cochlear implantation

Objectives: To investigate if the interaural time difference (ITD) ability is dependent of stimulation strategy. To examine the correlation between ITD, interaural level differences (ILD) and the ability to localize different sounds.

Methods: Thirty subjects aged 8–13 who were implanted bilaterally before 3 years of age were tested. Twenty of the subjects used processors programmed with fine structure (FS) strategy on both sides. ITD and ILD just noticeable difference (JND) of a 250?Hz pure tone was measured using their clinical processors. Furthermore, their ability to localize sound in the horizontal plane was measured using eye tracking.

Results: Ten of the 20 subjects with FS obtained an ITD threshold compared to none in the group without FS (0/10). ILD JND was correlated to localization ability of the broadband (BB) sound. Mean absolute error of the localization of a low-frequency (LF) sound was larger than that of a BB sound.

Conclusions: The ability to detect ITD was present only when the cochlear implant stimulation had FS. The LF sound was more difficult to localize than the BB sound and ITD ability of FS strategies did not affect the localization ability of either sound. A low ILD seems necessary to improve the localization ability.     

Mapping lateralization of click trains in younger and older populations

The main purpose of this study was to describe and compare lateralization of earphone-presented stimuli in younger and older individuals. Lateralization functions, relating perceived location to either interaural time differences (ITDs) or interaural level differences (ILDs) were determined for 78 subjects, aged 21-88 years, who responded by pressing one of nine keys to indicate the perceived location of the stimulus. All subjects were healthy, without any history of hearing loss or ear surgery and within the normal pure tone audiometric range for their age group. Interaural pure tone and click thresholds did not differ by more than 5 dB across ears. The ILD lateralization functions, ranging from 10 dB favoring the left ear to 10 dB favoring the right ear were linear. In contrast, the ITD lateralization functions were S-shaped with a clear linear component ranging from 750 micros favoring one ear to 750 micros favoring the other ear and with an asymptote from 750 micros to 1 ms. The same general shape of the ITD and ILD lateralization functions was found at all ages, but the linear slope of the ITD lateralization function became shallower with age. The ability to discriminate midline-located click trains (ITD and ILD=0) from ITD-lateralized click trains deteriorated with age, while the comparable ability to discriminate ILD-lateralized click trains did not change significantly with age. The data support two general conclusions. First there seems to be an overall reduction in the range of ITD-based lateralization due to aging. Second, there is a greater reduction in sensitivity due to aging in changes from the perceived midline position (ITD and ILD=0) when ITD is manipulated than when ILD is manipulated.     

Interaural Correlation Fails to Account for Detection in a Classic Binaural Task: Dynamic ITDs Dominate N0Sπ Detection

Binaural signal detection in an NoSπ task relies on interaural disparities introduced by adding an antiphasic signal to diotic noise. What metric of interaural disparity best predicts performance? Some models use interaural correlation; others differentiate between dynamic interaural time differences (ITDs) and interaural level differences (ILDs) of the effective stimulus. To examine the relative contributions of ITDs and ILDs in binaural detection, we developed a novel signal processing technique that selectively degrades different aspects (potential cues) of binaural stimuli (e.g., only ITDs are scrambled). Degrading a particular cue will affect performance only if that cue is relevant to the binaural processing underlying detection. This selective scrambling technique was applied to the stimuli of a classic N0Sπ task in which the listener had to detect an antiphasic 500-Hz signal in the presence of a diotic wideband noise masker. Data obtained from five listeners showed that (1) selective scrambling of ILDs had little effect on binaural detection, (2) selective scrambling of ITDs significantly degraded detection, and (3) combined scrambling of ILDs and ITDs had the same effect as exclusive scrambling of ITDs. Regarding the question which stimulus properties determine detection, we conclude that for this binaural task (1) dynamic ITDs dominate detection performance, (2) ILDs are largely irrelevant, and (3) interaural correlation of the stimulus is a poor predictor of detection. Two simple stimulus-based models that each reproduce all binaural aspects of the data quite well are described: (1) a single-parameter detection model using ITD variance as detection criterion and (2) a compressive transformation followed by a crosscorrelation analysis. The success of both of these contrasting models shows that our data alone cannot reveal the mechanisms underlying the dominance of ITD cues. The physiological implications of our findings are discussed.     

Modulation Enhancement in the Electrical Signal Improves Perception of Interaural Time Differences with Bimodal Stimulation

Interaural timing cues are important for sound source localization and for binaural unmasking of speech that is spatially separated from interfering sounds. Users of a cochlear implant (CI) with residual hearing in the non-implanted ear (bimodal listeners) can only make very limited use of interaural timing cues with their clinical devices. Previous studies showed that bimodal listeners can be sensitive to interaural time differences (ITDs) for simple single- and three-channel stimuli. The modulation enhancement strategy (MEnS) was developed to improve the ITD perception of bimodal listeners. It enhances temporal modulations on all stimulated electrodes, synchronously with modulations in the acoustic signal presented to the non-implanted ear, based on measurement of the amplitude peaks occurring at the rate of the fundamental frequency in voiced phonemes. In the first experiment, ITD detection thresholds were measured using the method of constant stimuli for five bimodal listeners for an artificial vowel, processed with either the advanced combination encoder (ACE) strategy or with MEnS. With MEnS, detection thresholds were significantly lower, and for four subjects well within the physically relevant range. In the second experiment, the extent of lateralization was measured in three subjects with both strategies, and ITD sensitivity was determined using an adaptive procedure. All subjects could lateralize sounds based on ITD and sensitivity was significantly better with MEnS than with ACE. The current results indicate that ITD cues can be provided to bimodal listeners with modified sound processing.     

Neural and Behavioral Sensitivity to Interaural Time Differences Using Amplitude Modulated Tones with Mismatched Carrier Frequencies

Bilateral cochlear implantation is intended to provide the advantages of binaural hearing, including sound localization and better speech recognition in noise. In most modern implants, temporal information is carried by the envelope of pulsatile stimulation, and thresholds to interaural time differences (ITDs) are generally high compared to those obtained in normal hearing observers. One factor thought to influence ITD sensitivity is the overlap of neural populations stimulated on each side. The present study investigated the effects of acoustically stimulating bilaterally mismatched neural populations in two related paradigms: rabbit neural recordings and human psychophysical testing. The neural coding of interaural envelope timing information was measured in recordings from neurons in the inferior colliculus of the unanesthetized rabbit. Binaural beat stimuli with a 1-Hz difference in modulation frequency were presented at the best modulation frequency and intensity as the carrier frequencies at each ear were varied. Some neurons encoded envelope ITDs with carrier frequency mismatches as great as several octaves. The synchronization strength was typically nonmonotonically related to intensity. Psychophysical data showed that human listeners could also make use of binaural envelope cues for carrier mismatches of up to 2–3 octaves. Thus, the physiological and psychophysical data were broadly consistent, and suggest that bilateral cochlear implants should provide information sufficient to detect envelope ITDs even in the face of bilateral mismatch in the neural populations responding to stimulation. However, the strongly nonmonotonic synchronization to envelope ITDs suggests that the limited dynamic range with electrical stimulation may be an important consideration for ITD encoding.     

Models of Brainstem Responses to Bilateral Electrical Stimulation

A simple, biophysically specified cell model is used to predict responses of binaurally sensitive neurons to patterns of input spikes that represent stimulation by acoustic and electric waveforms. Specifically, the effects of changes in parameters of input spike trains on model responses to interaural time difference (ITD) were studied for low-frequency periodic stimuli, with or without amplitude modulation. Simulations were limited to purely excitatory, bilaterally driven cell models with basic ionic currents and multiple input fibers. Parameters explored include average firing rate, synchrony index, modulation frequency, and latency dispersion of the input trains as well as the excitatory conductance and time constant of individual synapses in the cell model. Results are compared to physiological recordings from the inferior colliculus (IC) and discussed in terms of ITD-discrimination abilities of listeners with cochlear implants. Several empirically observed aspects of ITD sensitivity were simulated without evoking complex neural processing. Specifically, our results show saturation effects in rate–ITD curves, the absence of sustained responses to high-rate unmodulated pulse trains, the renewal of sensitivity to ITD in high-rate trains when inputs are amplitude-modulated, and interactions between envelope and fine-structure delays for some modulation frequencies.